Nervous System.—The nervous centre of the Rotifera is the brain (Fig. 112, C, br), a ganglion lying dorsal to the pharynx; and when this is short it may be immediately below the surface of the disc (Microcodon). In Bdelloida a second ganglion is present below the pharynx, and is connected with the former by lateral cords which contain ganglion cells. From the brain, nerves are given off to the disc, to the muscles, and to the integument of the body, as well as to the sense organs. The largest nerves are two given off from the sides of the brain, each of which divides into a lateral and a ventral trunk, which run nearly the whole length of the animal.
The brain of several Notommatidae has a curious appendage, white by reflected light and very opaque; it is a sac full of chalky mineral matter, which dissolves readily in dilute acids.
Sense Organs.—The most widely diffused sense organs are the antennae or feelers, which may serve for touch or smell, or possibly both. Each antenna is a conical or tubular outgrowth of the skin; from its apex projects a fine pencil of sense hairs borne on a protoplasmic cushion, which receives a nerve. Often the antenna is elongated, and may then contain a muscle by which it is retractile (lateral antennae of Melicerta); sometimes it is reduced to a slight prominence bearing the setae (dorsal antenna of this genus). There are usually three antennae—a median dorsal (Figs. 109, B, a, and 112, C, am) and two lateral (Figs. 106, 112, C, and 115, A, al), often approximated towards the ventral surface, and sometimes all but fused on the middle line, or completely united (Conochilus dossuarius, Copeus caudatus).[[259]]
Most Rotifers possess an organ of sight. This in its simplest form is a refractive globule seated in a red pigmented cup through which the nerve passes; in other cases it lies directly on the brain. Very frequently the eye is paired (Figs. 112, B, and 115, A); and these paired eyes may lie on the brain, and then are so close together that the pigment-cups have the shape of an x, or else they are seated in the dorsal region of the head behind the disc. In some cases they lie just under the ciliary wreath, or even within the region of the disc, and pass towards its ventral side in Pedalion (Fig. 117, A, e). In Rotifer they lie just under the dorsal side of the proboscis just below its apex. The median and two lateral eyes often exist together, as in Eosphora; and sometimes additional paired eyes exist. In Furcularia longiseta, var. grandis a pair of pigment spots (eyes?) occurs at the hinder end of the body just in front of the foot.
The active Ploima show a spontaneity of movement and marked power of avoiding obstacles, etc. This is still more marked in the very active Pedalion, which, as Rousselet notes, clearly avoids capture by the dropping tube, aided by its sense of sight, as he suggests, or by the tactile or olfactory powers of the antennae. They must rank as psychically high in the scale of creatures of simple organisation.
Reproductive Organs and Reproduction.—The most conspicuous organ in the female is the large yolk-gland or vitellarium (Figs. 106 and 109, A, vm), which was regarded as the ovary by all the older observers. It consists usually of eight cells, with conspicuous nuclei, lying on the ventral side of the stomach, and frequently displaced to one side; but in most Asplanchnidae it forms a broad transverse band of numerous cells. In Pterodina it is horseshoe-shaped, while in Seisonaceae and Bdelloida it is paired, either gland containing four or eight cells. The true ovary or germarium (Fig. 106, gm) lies more or less hidden between the yolk-gland and the stomach; it is composed of numerous minute rounded cells, of which the hindmost for the time being enlarges by nutrition from the yolk-gland, and finally receives a membranous shell. This true ovary is somewhat lateral in most Rotifers, but is median in Asplanchnidae, and paired in Pterodina, Bdelloida, and Seisonaceae. A membranous covering is common to the ovary and yolk-gland (paired when these are paired); it is continued into a thin-walled tube or oviduct, which opens into the cloaca on its ventral side beyond the bladder or common renal duct. In the viviparous species the mature ovum (Fig. 112, em) usually lies in the oviduct, dilating it into a sort of "uterus" until the birth of the young. The ordinary eggs or "summer eggs" are formed without any fertilisation, and develop immediately; they are often hatched within the tube of the tubicolous species.
Under certain conditions the unfertilised females produce exclusively smaller eggs, which develop into males. Maupas[[260]] has demonstrated that a rise in temperature to a minimum of 26° C. (79° F.) is the efficient factor. But as Bergendal points out,[[261]] the critical temperature probably varies with the antecedent conditions of the race, since males occur in Greenland at a very much lower temperature; and it would seem probable that a temperature approaching that at which the pools habitually dry up is what is necessary for the production of males, as a provision for those fertilised eggs, which, having a hard shell often adorned with prickly prominences, and usually remaining for some time before development, are capable of withstanding drought; such eggs are termed "winter eggs," but a better term would be "resting eggs" (German, "Dauereier").[[262]]
Fig. 113.—Diglena catellina. (After Weber.) A, Male; B, the pair in copula; C, female, p, Penis; te, testis.
The male organs consist of a testis (Fig. 113, A, te) with accessory glands, a large seminal vesicle, and a protrusible or projecting penis (p). In Notommata and Diglena true intromission at the cloaca (B) has been seen by many observers; but it appears equally certain that in many cases the male bores into the body-wall of the female at any point, and deposits the spermatozoa in the body-cavity, so that they must pass through the wall of the oviduct to effect fertilisation. Maupas finds that the process of fertilisation is ineffective except upon such newly-hatched females as would otherwise be the parents of small male eggs; that fertilisation is inoperative even for these at a later age when their eggs have begun to mature; and that it is wholly useless for those that lay ordinary summer eggs. The parent of male or winter eggs would thus be comparable to the queen bee, which if not fertilised produces drones. These sexual relations find a close parallel in the Ostracod and Phyllopod Crustacea, as well as in many plant-lice (Homoptera).